{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T05:02:18Z","timestamp":1787029338089,"version":"build-2736575974"},"reference-count":27,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2025,9,15]],"date-time":"2025-09-15T00:00:00Z","timestamp":1757894400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Hebei Key Laboratory of Floral Biological Breeding","award":["21326301D"],"award-info":[{"award-number":["21326301D"]}]},{"name":"Hebei Key Laboratory of Floral Biological Breeding","award":["KY2024058"],"award-info":[{"award-number":["KY2024058"]}]},{"name":"Hebei Key Laboratory of Floral Biological Breeding","award":["YJ2025010"],"award-info":[{"award-number":["YJ2025010"]}]},{"name":"Hebei Province Science and technology plan project","award":["21326301D"],"award-info":[{"award-number":["21326301D"]}]},{"name":"Hebei Province Science and technology plan project","award":["KY2024058"],"award-info":[{"award-number":["KY2024058"]}]},{"name":"Hebei Province Science and technology plan project","award":["YJ2025010"],"award-info":[{"award-number":["YJ2025010"]}]},{"name":"Basic Scientific Research Projects of Provincial Universities","award":["21326301D"],"award-info":[{"award-number":["21326301D"]}]},{"name":"Basic Scientific Research Projects of Provincial Universities","award":["KY2024058"],"award-info":[{"award-number":["KY2024058"]}]},{"name":"Basic Scientific Research Projects of Provincial Universities","award":["YJ2025010"],"award-info":[{"award-number":["YJ2025010"]}]},{"name":"Hebei Agricultural University","award":["21326301D"],"award-info":[{"award-number":["21326301D"]}]},{"name":"Hebei Agricultural University","award":["KY2024058"],"award-info":[{"award-number":["KY2024058"]}]},{"name":"Hebei Agricultural University","award":["YJ2025010"],"award-info":[{"award-number":["YJ2025010"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Ulmus pumila L. \u2018Zhonghua Jinye\u2019 is a plant variety with colourful leaves that is widely used in landscaping. In our study, the leaves of U. pumila \u2018Zhonghua Jinye\u2019 fade and turn green under light (LT, 45%), moderate (MD, 70%) and high (HG, 95%) shading treatment, reducing its ornamental value. However, the mechanism underlying this adaptation to shade is poorly understood. The objective of this study was to elucidate the mechanism of physiological, cellular microstructural and transcriptional changes involved in leaf regreening in U. pumila \u2018Zhonghua Jinye\u2019. Our results showed that the pigment content of U. pumila \u2018Zhonghua Jinye\u2019 leaves increased under shade stress, with a corresponding colour change from yellow to dark green. Thus, U. pumila \u2018Zhonghua Jinye\u2019 adapted to shade stress by increasing leaf pigment and chlorophyll content. Transmission electron microscopy showed that thylakoid stacking in the grana lamellae changed significantly from a loose state to a closely packed structure under shaded conditions. Because plant pigments were located mainly in thylakoids, this closer stacking increased photosynthetic efficiency and pigment accumulation. RNA sequencing analysis showed that Lhcb1, a key thylakoid membrane gene, was upregulated under shade, which promoted thylakoid stacking and light absorption. In the chlorophyll synthesis pathway, haeme metabolism was inhibited, increasing protoporphyrin IX flow to the chlorophyll pathway and promoting the synthesis of chlorophyll a\/b. The simultaneous upregulation of plant hormone-related genes promoted an increase in plant leaf area, improving the light energy utilisation ratio. This study is the first to report the self-regulatory mechanism that leads to colour change in U. pumila \u2018Zhonghua Jinye\u2019 under shade stress and provides a theoretical basis for the cultivation of tree species with colourful leaves.<\/jats:p>","DOI":"10.3390\/plants14182868","type":"journal-article","created":{"date-parts":[[2025,9,15]],"date-time":"2025-09-15T16:27:33Z","timestamp":1757953653000},"page":"2868","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Analysis of the Metabolic and Structural Changes in Ulmus pumila \u2018Zhonghua Jinye\u2019 Leaf Under Shade Stress"],"prefix":"10.3390","volume":"14","author":[{"given":"Yichao","family":"Liu","sequence":"first","affiliation":[{"name":"Hebei Forestry and Grassland Science Research Institute, Shijiazhuang 050000, China"},{"name":"College of Forestry College (Landscape and Tourism), Hebei Agricultural University, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6176-1925","authenticated-orcid":false,"given":"Yongtan","family":"Li","sequence":"additional","affiliation":[{"name":"College of Forestry College (Landscape and Tourism), Hebei Agricultural University, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ning","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Forestry College (Landscape and Tourism), Hebei Agricultural University, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lihui","family":"Zuo","sequence":"additional","affiliation":[{"name":"College of Landscape and Ecological Engineering, Hebei University of Engineering, Handan 056000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yang","family":"Zhou","sequence":"additional","affiliation":[{"name":"College of Forestry College (Landscape and Tourism), Hebei Agricultural University, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8487-5896","authenticated-orcid":false,"given":"Ping","family":"Li","sequence":"additional","affiliation":[{"name":"College of Forestry College (Landscape and Tourism), Hebei Agricultural University, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shijie","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Landscape and Ecological Engineering, Hebei University of Engineering, Handan 056000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shuxiang","family":"Feng","sequence":"additional","affiliation":[{"name":"Hebei Forestry and Grassland Science Research Institute, Shijiazhuang 050000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shufang","family":"Yan","sequence":"additional","affiliation":[{"name":"Hebei Forestry and Grassland Science Research Institute, Shijiazhuang 050000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yinran","family":"Huang","sequence":"additional","affiliation":[{"name":"College of Forestry College (Landscape and Tourism), Hebei Agricultural University, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Minsheng","family":"Yang","sequence":"additional","affiliation":[{"name":"College of Forestry College (Landscape and Tourism), Hebei Agricultural University, Baoding 071000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,9,15]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1007\/s11240-007-9310-8","article-title":"A protocol for Ulmus minor Mill. micropropagation and acclimatization","volume":"92","author":"Conde","year":"2008","journal-title":"Plant Cell Tissue Organ Cult."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"1281","DOI":"10.1007\/s11738-012-1167-8","article-title":"Cadmium toxicity affects photosynthesis and plant growth at different levels","volume":"35","author":"Dias","year":"2013","journal-title":"Acta Physiol. Plant."},{"key":"ref_3","first-page":"112","article-title":"Caryopteris \u00d7 clandonensis \u2018worcester gold\u2019 and its domestication, culture","volume":"31","author":"Yuan","year":"2004","journal-title":"Acta Hortic. Sin."},{"key":"ref_4","first-page":"352","article-title":"Progress on color emerging mechanism and photosynthetic characteristics of colored-leaf plants","volume":"32","author":"Jiang","year":"2005","journal-title":"Acta Hortic. Sin."},{"key":"ref_5","first-page":"717","article-title":"Response of pigment content of golden-leaf plants to light intensity","volume":"34","author":"Hu","year":"2007","journal-title":"Acta Hortic. Sin."},{"key":"ref_6","first-page":"591","article-title":"Contents of pigments and anatomical structure of leaves in Acer negundo \u2018Aurea\u2019","volume":"43","author":"Kang","year":"2023","journal-title":"Bull. Bot. Res."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"64","DOI":"10.1007\/s11295-017-1139-7","article-title":"Transcriptome analysis of Ulmus pumila \u2018Jinye\u2019 responses to different shading involved in chlorophyll metabolism","volume":"13","author":"Zhang","year":"2017","journal-title":"Tree Genet. Genomes"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Zuo, L., Zhang, S., Liu, Y., Huang, Y., Yang, M., and Wang, J. (2019). The reason for growth inhibition of Ulmus pumila \u2018Jinye\u2019: Lower resistance and abnormal development of chloroplasts slow down the accumulation of Energy. Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20174227"},{"key":"ref_9","first-page":"9","article-title":"The effects of different light treatments on the leaf color and physiological indicators of Berberis thunbergii var","volume":"25","author":"Chen","year":"2019","journal-title":"Atropurpurea. Anhui Agri. Sci. Bull."},{"key":"ref_10","unstructured":"Zou, Q. (2000). Experimental Instruction of Plant Physiology, China Agricultural Press."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1035","DOI":"10.1093\/bioinformatics\/btt087","article-title":"EBSeq: An empirical bayes hierarchical model for inference in RNA-seq experiments","volume":"29","author":"Leng","year":"2013","journal-title":"Bioinformatics"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1038\/75556","article-title":"Gene ontology: Tool for the unification of biology","volume":"25","author":"Ashburner","year":"2000","journal-title":"Nat. Genet."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1093\/nar\/28.1.33","article-title":"The COG database: A tool for genome scale analysis of protein functions and evolution","volume":"28","author":"Tatusov","year":"2000","journal-title":"Nucleic Acids Res."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"D115","DOI":"10.1093\/nar\/gkh131","article-title":"UniProt: The universal protein knowledgebase","volume":"32","author":"Apweiler","year":"2004","journal-title":"Nucleic Acids Res."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1093\/nar\/gkh063","article-title":"The KEGG resource for deciphering the genome","volume":"32","author":"Kanehisa","year":"2004","journal-title":"Nucleic Acids Res."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Koonin, E.V., Fedorova, N.D., Jackson, J.D., Jacobs, A.R., Krylov, D.M., Makarova, K.S., Mazumder, R., Mekhedov, S.L., Nikolskaya, A.N., and Rao, B.S. (2004). A comprehensive evolutionary classification of proteins encoded in complete eukaryotic genomes. Genome Biol., 5.","DOI":"10.1186\/gb-2004-5-2-r7"},{"key":"ref_17","first-page":"71","article-title":"Integrated nr database in protein annotation system and its localization","volume":"32","author":"Deng","year":"2006","journal-title":"Comput. Eng."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"316","DOI":"10.1093\/nar\/gkr483","article-title":"KOBAS 2.0: A web server for annotation and identification of enriched pathways and diseases","volume":"39","author":"Xie","year":"2011","journal-title":"Nucleic Acids Res."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"755","DOI":"10.1093\/bioinformatics\/14.9.755","article-title":"Profile hidden markov models","volume":"14","author":"Eddy","year":"1998","journal-title":"Bioinformatics"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Jason, E., and Ziv, B.J. (2006). STEM: A tool for the analysis of short time series gene expression data. BMC Bioinform., 7.","DOI":"10.1186\/1471-2105-7-191"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"879","DOI":"10.1007\/s11033-008-9258-8","article-title":"Identification and characterization of class 1 DXS gene encoding 1-deoxy-D-xylulose-5-phosphate synthase, the first committed enzyme of the MEP pathway from soybean","volume":"36","author":"Zhang","year":"2009","journal-title":"Mol. Biol. Rep."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1146\/annurev.pp.26.060175.001015","article-title":"Chlorophyll proteins: Light-harvesting and reaction center components of plants","volume":"26","author":"Thornber","year":"1975","journal-title":"Annu. Rev. Plant Physiol."},{"key":"ref_23","first-page":"250","article-title":"Structure and function of chloroplast membranes: Effect of potassium and magnesium lons on the strcture and absorption spectrum of two kinds chloroplast membranes","volume":"21","author":"Zhang","year":"1980","journal-title":"Acta Bot. Sin."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1007\/s11120-016-0252-z","article-title":"Light acclimation of shade-tolerant and light-resistant tradescantia species: Induction of chlorophyll a fluorescence and p700 photooxidation, expression of psbs and lhcb1 proteins","volume":"130","author":"Mishanin","year":"2016","journal-title":"Photosynth. Res."},{"key":"ref_25","unstructured":"Raghavendra, A.S. (1998). Light-harvesting complexes of higher plants. Photosynthesis: A Comprehensive Treatise, Cambridge University Press."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1007\/s11103-004-1963-7","article-title":"Constitutive expression of pea Lhcb 1-2 in tobacco affects plant development, morphology and photosynthetic capacity","volume":"55","author":"Labate","year":"2004","journal-title":"Plant Mol. Biol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"3646","DOI":"10.1105\/tpc.114.127373","article-title":"The light-harvesting chlorophyll a\/b binding proteins Lhcb1 and Lhcb2 play complementary roles during state transitions in Arabidopsis","volume":"26","author":"Pietrzykowska","year":"2014","journal-title":"Plant Cell"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/14\/18\/2868\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,9]],"date-time":"2025-10-09T18:46:03Z","timestamp":1760035563000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/14\/18\/2868"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,9,15]]},"references-count":27,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2025,9]]}},"alternative-id":["plants14182868"],"URL":"https:\/\/doi.org\/10.3390\/plants14182868","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,9,15]]}}}